L[Ev] A[Lexandrovitch] Nessov DINOSAURS of NORTHERN EURASIA: NEW DATA ABOUT ASSEMBLAGES, ECOLOGY, and PALEOBIOGEOGRAPHY

Saint Petersburg University Institute of Earth Crust L[ev] A[lexandrovitch] Nessov DINOSAURS OF NORTHERN EURASIA: NEW DATA ABOUT ASSEMBLAGES, ECOLOGY, AND PALEOBIOGEOGRAPHY Reviewers: Corresponding Member of the Russian Academy of Sciences I. S. Darevsky Doctor of Geological and Mathematical Sciences V. A. Prozorovsky Printed by resolution of the Editorial Council, Geology Faculty, St. Petersburg State University St. Petersburg, 1995 Translator: Tatayana Platonova, Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia Translation editor: Hans-Dieter Sues, Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A. NOTE ON TRANSLATION: This work, originally published in Russian in 1995, has been translated into English to introduce Nessov’s research on dinosaurs from the former Soviet Union to specialists worldwide. It contains important primary descriptions for a number of new dinosaurian taxa from the Cretaceous of Uzbekistan as well as a comprehensive synopsis of localities with dinosaurian remains on the territory of the former Soviet Union. Some of the paleobiological speculations will strike western readers as idiosyncratic, but we tried to retain the original character of Nessov’s work as much as possible in this translation. Throughout the text, pagination for the Russian original is indicated by bold-face numbers in brackets to facilitate cross-referencing. This translation should be used in conjunction with the original work to view the illustrations (which are not reproduced here). The transliteration used by the translator differs in some respects from that commonly used in the United States so that the spelling of certain names of authors and places differ (e.g., “Yefremov” rather than “Efremov”). Ms. Platonova’s translation effort was supported by National Science Foundation grant EAR-9804771. TABLE OF CONTENTS [WITH ORIGINAL PAGINATION] Introduction 4 1. Review of major dinosaurian taxa of the Middle Jurassic-Cretaceous of northern Eurasia 7 1.1.Segnosaurs 7 1.2.Theropods 15 1.3.Sauropods 19 1.4. Stegosaurs and ankylosaurs 20 1.5. Ornithopods 21 1.6. Ceratopsia 25 2. Descriptions of new discoveries 35 3. Some features of the habitat and dispersal routes of dinosaurs 58 4. Appendix. Localities of remains of bones, teeth, eggshell and tracks of dinosaurs in northern Eurasia 78 4.1. Bones and teeth of dinosaurs in Mesozoic deposits 80 4.1.1. Jurassic 80 4.1.2. Lower Cretaceous 84 4.1.3. Upper Cretaceous 89 4.2. Remains of Cretaceous dinosaurs redeposited in Paleogene sediments 112 4.3. Eggs, clutches, and eggshell of dinosaurs from the Cretaceous 113 4.4. Tracks of dinosaurs 115 4.4.1. Jurassic 115 4.4.2. Cretaceous 118 4.5. Remains that should no longer be attributed to dinosaurs and localities that should no longer be regarded as those where of dinosaurian remains were found in the CIS 120 References 125 Summary 144 Legends for plates 146 Index of taxonomic names of dinosaurs of family rank and lower, discovered within the boundaries of the former Soviet Union 152 Index of localities 154 INTRODUCTION The present work is devoted to the dinosaurs of northern Eurasia within the boundaries of the former Soviet Union. Remains and tracks of dinosaurs are known from Uzbekistan, Kazakhstan, Tadzhikistan, Kirghizia, Turkmenistan, Russia, Armenia, and Georgia. Existing reviews of localities of dinosaurian remains discovered in the former Soviet Union (Tolmachev, 1926; Yuryev, 1954a) as well as specialized bibliographical selections of discoveries of these animals in that country (Riabinin, 1931a, 1939; Rozhdestvensky, 1964; Rozhdestvensky and Tatarinov, 1964: p. 647, 648) usually include, in addition to dinosaurs, other ancient reptiles from either particular regions or from all of Asia (Yefremov, 1944; Rozhdestvensky, 1964, 1974; Rozhdestvensky, 1977; Rozhdestvensky and Khozatsky, 1967). Some reviews are incomplete (Rozhdestvensky and Tatarinov, 1964; Paleozoology of the USSR). All have become obsolete. If considered from the perspective of cladistic systematics, dinosaurs are a paraphyletic group because they do not include the ancestor of the group nor all its descendants (as birds are the descendants of theropod dinosaurs). Dinosaurs, whether they represent one, two, or possibly several lineages, take their origin from pseudosuchians. Therefore, it would be more natural, from the cladistic point of view, to establish a group including various pseudosuchians, pterosaurs(?), as well as crocodylians, ornithischians, and saurischians, and also birds or a group including only several evolutionarily derived theropods and birds. It would nevertheless be reasonable to preserve the term "dinosaur" in order to be free to use it in the future. The first discoveries and/or identifications of dinosaurian remains in the European part of the Commonwealth of Independent States (CIS) were made by Kiprianov prior to 1883, K. F. Egorov in 1912 and P. M. Klevensky in 1931, by A. D. Arkhangelsky in Transbaikalia in 1910, by V. T. Surgai and A. F. Sosedko in the Kyzylkum in 1930, by E. V. Ivanov in 1923 and V. D. Prinada in Tashkent Chul' [chul' = hilly plain; Tashkent Chul' is a hilly plain located north of Tashkent in southern Kazakhstan] in 1924-1925, by P. D. Trusov in 1928 and O. S. Vyalov in the Fergana region in the 1930's, by G. V. Mukhin in 1925, A. N. Kirikov in 1926, E. I. Belyaeva and M. G. Prokhorov in 1927 and D. I. Yakovlev in the basin of the Il River, and by I. I. Krom in the basin of the Emba River in 1926. Shell of dinosaurian eggs was found for the first time in the former Soviet Union by V. S. Erofeyev and L. D. Kudina in the Zaisan Basin in 1959. Tracks of dinosaurs were first discovered on the conterminous territory of the CIS by G. D. Romanovsky and P. A. Chuenko in 1882, and tracks of dinosaurs were found by Chabukiani in Georgia in 1933. Citizens of Russia (inhabitants of the Kasatkina stanitsa [stanitsa = large cossack village], now the village of Kasatkino) and later Colonel Manakin discovered, for the first time for science, a locality with dinosaurian remains on Chinese territory (1900-1902) (Manakin, 1902; Riabinin, 1925). This find was made long before the discovery of Central Asian localities (in Mongolia and also the Chinese provinces of Gansu and Inner Mongolia). Over the past 15 to 20 years, the amount of data on the composition of dinosaurian faunas in this country [Commonwealth of Independent States, abbreviated as CIS] and the list of localities have increased several times. [4/5] Therefore, a new review of the evidence is needed. Ichnological research on dinosaurs is proceeding vigorously, with studies being carried out by V. V. Kurbatov, V. P. Novikov, K. N. Amanniyazov, V. I. Sedletsky, and L. K. Gabuniya. These efforts have also been guided by results of paleoichnology abroad (Lockley, 1991; Lockley et al., 1991). A great body of new data was obtained by teams of the Stratigraphic Party headed by I. M. Abduazimova (Tashkent) and Leningrad (St. Petersburg) University. It was not uncommon that localities, where previously only fragmentary remains of one or two dinosaurian forms had been recorded, appeared to contain rich assemblages of these animals after several years of work. The Appendix to the present work lists all known localities, including even those that contain few dinosaurian remains that cannot be identified even to the level of order. Several of these localities may represent the first evidence of rich and previously unknown dinosaurian faunal complexes. Thus, for instance, even after dedicated paleontological exploration at Dzharakuduk, the fauna from this locality was regarded as relatively impoverished (Rozhdestvensky, 1964). Now, however, the locality is known to contain the most diverse fauna of aquatic and terrestrial vertebrates from the Cretaceous of Eurasia. It should be noted that very little is known in the world as a whole and in Eurasia in particular about the dinosaurs during the period from the end of the Middle Jurassic to the beginning of the Late Jurassic (Bathonian-Callovian; Oxford Clay) and particularly about the faunas of these animals during the first half of the Late Cretaceous (Cenomanian - Santonian) (Steel, 1969, 1970; Haubold, 1979; Steel and Haubold, 1979; Norman, 1985; Czerkas and Olson, 1987; Chure and McIntosh, 1989; Russell and Weishampel, 1989; Norman, 1989; Weishampel et al., 1990b; Britt et al., 1990). Even already available data on the dinosaurs of that time given in the literature are very incomplete (Charig, 1993: fig. 3). Even fewer data on dinosaurs are available for the Aalenian stage (at the beginning of the Middle Jurassic). No significant remains of these animals are known worldwide for this age interval. During the past few years, remains of dinosaurs have been found on the territory of the former Soviet Union that belong to taxa previously unknown for this region: Segnosauria and Oviraptorosauria in the Cretaceous, Stegosauria in the Jurassic, and also true horned dinosaurs, Ceratopsidae, for which no significant records had been known for the Cretaceous of Eurasia and which had long been considered endemic to North America (Rozhdestvensky, 1977; Coombs, 1984; Russell, 1991). The world's oldest, or at least very early, representatives of several taxa were found: Hadrosauridae in the late Albian, Protoceratopsidae in the late Albian and early Cenomanian, Ceratopsidae in the late Turonian and Coniacian, Segnosauria in the Albian, early Cenomanian, and possibly late Barremian - middle Aptian, Caenagnathidae in the late Turonian, the genus Troodon in the early Cenomanian, and Stegosauria at the boundary between the Middle and Late Jurassic. All these data as well as other new evidence significantly clarify possible dispersal routes, evolutionary histories of taxa, and the evolutionary history of some dinosaurian assemblages from the Central Asian part of the CIS, Kazakhstan, Transbaikalia, the Far East and North-East of Russia, and also China, Mongolia, and western North America. Unfortunately, only a few complete skeletons of dinosaurs have been found on the territory of the former Soviet Union.

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